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I Taxonomic Review of Rhinolophus Pusillus and Rhinolophus Lepidus

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I Taxonomic Review of Rhinolophus Pusillus and Rhinolophus Lepidus Taxonomic Review of Rhinolophus pusillus and Rhinolophus lepidus (Chiroptera: Rhinolophidae) in Thailand Ariya Dejtaradol A Thesis Submitted in Partial Fulfillment of the Requirements for the Degree of Master of Science in Ecology (International Program) Prince of Songkla University 2009 Copyright of Prince of Songkla University i Thesis Title A Taxonomic Review of Rhinolophus pusillus and Rhinolophus lepidus (Chiroptera: Rhinolophidae) in Thailand Author Miss Ariya Dejtaradol Major Program Ecology (International Program) Major Advisor Examining Committee: ………………………………………. .....………….………………Chairperson (Asst. Prof. Dr. Sara Bumrungsri) (Assoc. Prof. Dr. Kitichate Sridith) .……..…….………………………..…… Co-advisor (Dr. Chavalit Vidthayanon) ………………………………………. .……..…….………………………..…… (Assoc. Prof. Dr. Chutamas Satasook) (Asst. Prof. Dr. Sara Bumrungsri) ………………………………………. .……..…….………………………..…… (Dr. Paul J. J. Bates) (Assoc. Prof. Dr. Chutamas Satasook) The Graduate School, Prince of Songkla University, has approved this thesis as partial fulfillment of the requirements for Master of Science Degree in Ecology (International Program) ………………………………………... (Assoc. Prof. Dr. Krerkchai Thongnoo) Dean of Graduate School ii กกกกก () กก 2551 กก (Rhinolophus pusillus ) กก (R. lepidus) กกก ก กก กกกก กก กกก 2 (synonyms ) ( subspecies ) กก กกกก กกก กกก กก กกก ก กกกกกกก ก กก 3 ก ก ก ก (Rhinolophus pusillus ) ก (Rhinolophus lepidus ) Rhinolophus lepidus refulgens ก (Rhinolophus sp. ) ก กก ก กก 4 ก ก ก (Rhinolophus sp. ) ก กกก ก กกก 2 กก กกกก iii Thesis Title A Taxonomic Review of Rhinolophus pusillus and Rhinolophus lepidus (Chiroptera: Rhinolophidae) in Thailand Author Miss Ariya Dejtaradol Major Program Ecology (International Program) Academic Year 2008 ABSTRACT Rhinolophus pusillus and R. lepidus are horseshoe bats which have essentially similar external morphology and cranio-dental measurements. Previous taxonomic studies suggested that their taxonomy was confusing as there were number of synonyms and a smaller number of subspecies. In the past, taxonomic studies of the two species based on both qualitative and quantitative characters of external and cranio-dental morphology. Today, morphometric analysis techniques which are more sophisticated and provide better result are commonly applied to discriminate between size-overlapped taxa. Currently, species identification can also be supported with analysis of echolocation calls. From the examination of bats in R. pusillus group throughout Thailand, there are possibly at least 3 taxa within this rhinolophid group based on morphological characters: R. pusillus , R. lepidus which is referred to subspecies R. l. refulgens that found in southern Thailand, and Rhinolophus sp. from Phu Suan Sai National Park, Loei Province which is significantly different from other taxa in this group. However, they can be divided into four groups based on echolocation frequency and morphometrics: R. l. refulgens , R. pusillus , Rhinolophus sp. from Loei Province and Rhinolophus sp. from Khao Samo Khon, Lopburi Province which has highest frequency of echolocation calls among the bats in this group. With the current evidences, it indicated the presence of the species complex in R. pusillus group, and there are more than two taxa of this group in Thailand. The further molecular study maybe warrant for taxonomic classification of these bats. iv ACKNOWLEDGEMENTS I am deeply indebted to Assistant Professor Dr. Sara Bumrungsri and Associate Professor Dr. Chutamas Satasook for their advice and recommendation. I am greatly indebted to Dr. Paul J. J. Bates of the Harrison Institute, UK, for his help and outstanding supervision. I would also like to thank Dr. David Harrison and Malcolm Pearch of the Harrison Institute for their help and advice. I am also grateful to the Thesis Examining Committees, including Associate Professor Dr. Kitichate Sridith of Department of Biology, Faculty of Science, Prince of Songkla University, and Dr. Chavalit Vidthayanon of World Wildlife Fund-Thailand, for their correction and valuable suggestion. This work was supported by Darwin Initiative (DEFRA) of the UK Government, the TRF/BIOTEC Special Program for Biodiversity Research and Training grant BRT T_151001, and Graduate school of Prince of Songkla University (PSU). I would like to thank Director-general of the Department of National Park, Wildlife and Plant Conservation (DNP) for research permission, to Prateep Rojanadilok, Siriporn Thong-Aree and Saksit Simcharoen, the officers of DNP for their kind cooperation. My thanks are also due to Dr. Surachit Waengsothorn of Thailand Institute of Scientific and Technological Research (TISTR) who permits me loan specimens from TISTR . I also thank Bounsavane Douangboubpha, Phansamai Phommexay, Phouthone Kingsada, Pipat Soisook, Tuanjit Srithongchuay, and all students at Bat Research Unit for their help in fieldwork and specimens preparation. I would like to thanks Piyathip Piyapan, Medhi Yokubol, Sunate Karapan and Kwan Nualcharoen for their advice and generous help. Elsewhere, I would like to thank Sebastien Puechmaille of University College Dublin for his advice. I also thank Beatrix Lanzinger for generous help and encouragement when I stayed at Harrison Institute, UK. I also thank Phannee Sa-ardrit, Bongkot Wichachucherd and Piyapong Petchabun and all staffs of the Princess Maha Chakri Sirindhorn Natural History Museum for their kindly help during a long working with the collections in the museum. I am also grateful to Paula Jenkins and all staff of the mammal department v and the libraries of The Natural History Museum, London for their help. I am grateful to all lecturers and staffs of Department of Biology, Faculty of Science, Prince of Songkla University for generous help. Thanks are also due to Vu Dinh Thong and Pham Duc Tien of Institute of Ecology and Biological Resources (IEBR), Vietnam for their specimens deposited in the Harrison Zoological Museum. I would like to thank Neil Furey for echolocation data of Rhinolophus pusillus from Vietnam. I also thank Charles M. Francis for the molecular data from Bats of Southeast Asia Project. Finally, I would like to thank my father, my mother and my family for their support and encouragement. Ariya Dejtaradol vi Contents Page Abstract iii Acknowledgements v Contents vii List of Tables viii List of Figures ix Chapter 1: Introduction 1 Chapter 2: Literature Review 4 Chapter 3: Material and Methods 20 3.1 Specimen examined 20 3.2 Study sites 20 3.3 Field Methods 28 3.4 Laboratory methods 37 3.5 Analysis 40 Chapter 4: Results 43 4.1 Morphology comparison with other taxa 43 4.2 Variations 48 4.3 Echolocation 54 4.4 Measurements 60 4.5 Systematics summary 80 Chapter 5: Discussion 94 Chapter 6: Conclusion 99 References 101 Appendixes 110 Vitae 121 vii List of Tables Page Table 1. Descriptive statistics for echolocation call characters of 4 populations 56 of Rhinolophus . Table 2. Descriptive statistics and T-test p-value of six echolocation call 57 characters in male and female of three populations of Rhinolophus in Thailand. Table 3. List of measurements and their abbreviations. 61 Table 4. Descriptive statistics for morphological measurements of Rhinolophus 62 spp.from Thailand, R. lepidus , R. l. refulgens and R. pusillus from type localities. Table 5. Descriptive statistics for external measurements of 4 populations of 64 Rhinolophus spp. in Thailand. Table 6. Descriptive statistics (Mean ± SD; mm) for external and cranial 66 measurements from 3 populations of Rhinolophus spp. in Thailand and T-test between male and female. Table 7. One-way ANOVA of the measurements in 14 characters that used in 69 PCA between 4 populations. Table 8. The result from Tukey HSD post hoc test for the measurements in 14 70 characters that used in PCA. viii List of Figures Page Figure 1. Front view of noseleaf of Rhinolophus lepidus 7 Figure 2. Lateral view of noseleaf of Rhinolophus lepidus 7 Figure 3. Distribution of R. lepidus from literature review. 16 Figure 4. Distribution of R. pusillus from literature review. 16 Figure 5. Type of echolocation used by bats. 18 Figure 6. The concrete conduit near headquarter of Chiang Dao Wildlife 25 Sanctuary, Chiang Mai which the bats live in. Figure 7. The nature trail in Phu Suan Sai National Park which surrounded 25 by hill evergreen forest with banana trees. Figure 8. The harp trap was set across the nature trail in Phu Pha San, Phu Si- 26 Than Wildlife Sanctuary which surrounded by mixed deciduous forest. Figure 9. The harp trap was set across the stream near the road to Ao Son, 26 Tarutao National Park which surrounded by lowland evergreen forest. Figure 10. Khao Samo Khon limestone outcrop which covered by mixed 27 deciduous forest and surrounded by paddy field. Figure 11. Rubber and oil palm plantation around Silawan limestone outcrop, 27 Patiew District, Chumphon Province. Figure 12. Field data sheet 32 Figure 13. Right wing, head and some measurements of Rhinolophus l. refulgens 32 Figure 14. Pettersson ultrasound detector D 240x in time-expansion (10x) mode, 34 connected to sound recorder i-RIVER model: iHP-120 Multi-codec jukebox Figure 15. Label for wet specimen and skull: front of the wet specimen label 35 and back of the wet specimen label. Figure 16. Dorsal view of the skull of Rhinolophus pusillus 39 Figure 17. Ventral view of the skull of Rhinolophus pusillus 39 Figure 18. Lateral view of the skull of Rhinolophus pusillus 39 Figure 19. R. l. refulgens call and the BatSound software cursors and 42 measurement point used for parameter measurements. ix List of Figures (Continued) Page Figure 20. The skulls in lateral view of a) syntype of R. minor ( pusillus ) 45 Horsfield, 1823 (BM(NH)1879.11.21.684) from Java, Indonesia; b) Type specimen of R. gracilis (R. pusillus ) ( ♀ BM(NH) 61.1747) from West Java, Indonesia; c) R. pusillus ( ♂ BM(NH) 10.4.7.8) from East Madura, Malaysia; d) R. pusillus (♂ PSUZC-MM07.264) from Mukdaharn, Northeastern Thailand. Figure 21. The skulls in lateral view of a) Holotype of R. monticola (R. l. lepidus ) 46 ( ♂ BM(NH) 1879.11.21.151) from Masuri, Pakistan; b) R. l. lepidus (♂ HZM 19.28161) from New Delhi, India; c) R.
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